Paypayrola panamensis H. E. Ballard, 2022

Ballard, Harvey E., 2022, A new Paypayrola (Violaceae) from Panama, with notes about the species complexes for the genus, Phytotaxa 542 (3), pp. 283-292 : 288-289

Paypayrola panamensis H. E. Ballard , sp. nov. ( Figs. 1–3 )

Type: — PANAMA. Panamá: El Llano-Cartí Road, Km 26.5, 9°19’N, 78°55’W, 200 m, Tree 8 m, petals yellow, 14 April 1985, de Nevers, Herrera & Charnley 5363 (holotype, here designated: MO4998611 !; isotypes: U1757190 !, US 3556948!, WIS!) .

Paypayrola panamensis differs most obviously from P. guianensis in its proportionally longer leaf blades, subsessile to short-pedunculate, short inflorescences, and puberulent sepals that are commonly slightly to distinctly longer than broad; from P. hulkiana in its proportionally broader leaf blades, proportionally broader elliptical to suborbicular sepals commonly with two (rarely four) distinct lateral nerves in addition to the medial nerve and a frequently truncate or retuse apex; and from both in its larger leaf blades with more lateral nerves, blade apex weakly subattenuate to narrowly rounded, and a densely flowered inflorescence.

Small trees, (1.5) 3–10 m tall. Stem glabrous throughout, bark gray to brownish-gray, branches often minutely longitudinally ridged. Stipules promptly caducous, narrowly attenuate-triangular to lance-attenuate, 6–8 mm long × 2 mm wide at the base, sometimes carinate, apex acuminate to attenuate. Leaves simple, alternate below, sub-opposite or clustered near the branch tips; petiole slightly swollen, on young leaves blackish and smooth, on older becoming woody, grayish-brown and transversely rugose or furrowed, (4)6–21 × 1.6–3.9 (medial) mm; leaf blade oblanceolate to broadly oblanceolate, 18.4–29.3 × (4.1)5.9–9.8, 2.6–3.9(4.6), subcoriaceous, drying dull gray-green above, distinctly paler below, base narrowly cuneate to cuneate or weakly convexly tapering, 26–74(83)°, apex tapering or subattenuate to a narrowly rounded tip, rarely convexly rounded to an obtuse or broadly rounded tip, 39–78(149)°, complete veins 8–12 on each side, margins entire. Inflorescences cauliflorous and ramiflorous, 1–4 unbranched and/or branched botryoids per node, inflorescence bracts caducous, rachis 7–20 × 1.0– 1.6 mm, subsessile to short-pedunculate, peduncle 0.6–3.8 mm, rachis densely flowered, at least distally. Mature flowers 6–15 per botryoid, the proximal commonly aborted; pedicel 0.6–1.9 × 0.8–1.2 mm, articulate, pedicel bracteoles ovate-lanceolate to broadly deltate-triangular, 0.9–1.3 × 0.5–1.3 mm; flower buds yellow in life, oblanceoloid, flowers, at full anthesis, 7–18 mm long. Sepals 5, quincuncial, subequal, outer shorter than inner, fleshy basally, narrowly to broadly oblong-elliptical or suborbicular, as long as to distinctly longer than broad, 1.6–3.9 × 1.4–2.7 mm, length: width ratio 1.0–1.7(2.0), base abruptly rounded to subtruncate, apex broadly rounded or frequently truncate to retuse, minutely puberulent adaxially, margins ciliolate, (1)3 or 5 nerves, frequently raised and prominent, small black subapical subconical to cylindrical appendage frequently terminating medial nerve. Corolla weakly bilaterally symmetrical, petals 5, quincuncial, bright yellow, fleshy, strongly recurved in life, one petal ca. 2 × broader in distal half than others, all petals with five obscure nerves visible upon hydration, glabrous or sparsely to moderately minutely puberulent adaxially in basal half, sometimes sparsely minutely hirtellous along costa, margins eciliate or minutely ciliolate, narrow petals weakly pandurate, convex in cross-section, apex narrowly rounded, 7–11 mm long × 1.4–2.2 mm wide in their distal half, broadest petal pandurate, v-shaped in cross-section, apex retuse, 10–17 mm long × 3.2–3.4 mm wide in its distal half. Androecium 1.6–2.6 mm tall, stamens 5, filaments fused into a 0.8–1.6 mm tall tube (measured to highest point between the stamens), tube glabrous or sparsely pilosulous, dorsal connective slightly exceeded by the thecae, connective appendage small, 0.14–0.27 mm tall, conical to narrowly triangular proximally and attenuate distally, projecting slightly beyond thecae, thecae slanting diagonally to connective appendage. Gynoecium 3.8–4.0 mm tall, pistil tricarpellate, compound, placentation parietal, ovary densely appressed-pilosulous, style glabrous, filiform, often bent in the middle, 2.7–2.9 mm long, stigma trilobate. Fruit a 3-valved capsule, ovoid to rhombic-ovoid, immature whitish to pale green in life, at maturity pale green or green in life, dried often with small irregular reddish streaks or blotches, smooth, glabrous or sparsely and minutely puberulent in basal one-third, 22–40 × 16–20 mm. Seeds 2–3 per carpel, oblong-ellipsoid to obovoid with broadly rounded apex, medium to dark reddish-brown, smooth and shiny, 8.7–9.5 × 5.1–5.8 mm including basal appendage.

Etymology:— Being the sole Paypayrola in Mesoamerica and well represented by many collections in a bimodal distribution of west-central and east-central Panama, the species is named for the country of Panama to which it is endemic.

Distribution and habitat:— Paypayrola panamensis is presently confirmed from 48 herbarium specimens representing 26 separate localities inferred as distinct populations in central Panama ( Fig. 3 ). The species grows in moist to wet evergreen forest on low slopes and low local ridgetops often near or along streams and rivers but rarely in riparian forest, on the Atlantic slope, to the west and east of the Canal Zone but not near it, at 30–500 m in elevation.

Phenology:— The species is in flower throughout the year. Fruits apparently mature within a month or two of flowering. Flower buds, mature open flowers, young fruits, and ripening or mature fruits are apparently produced on the same tree at all times of the year.

Proposed Conservation Status:— Conservation assessment using GeoCAT provided a “VU” (Vulnerable) designation based on Extent of Occurrence (5,852.847 km 2), and a “EN” (Endangered) designation based on Area of Occupancy (84.000 km 2). Endangered status is proposed as the most conservative conservation status for this rather narrow endemic.

Discussion:— Comparisons of herbarium specimens from Panama with those of South America, and studies of types and protologues for known species, have revealed many misidentifications. Correctly assigned specimens demonstrate that inflorescence morphology, corolla color, sepal shape, and to a lesser degree, leaf morphology, are fixed in individual species. Importantly, species are readily placed into three species complexes based on corolla color, with supporting traits of inflorescence and sepal morphology. Thus, the genus consists currently of eight species divided in three morphological groups. The Blanchetiana species complex possesses white, commonly pink-tinged, corollas, sepals much longer than broad, and inflorescences subsessile or short-pedunculate and densely flowered. This group consists of P. arenacea , narrowly endemic to Amazonian Venezuela; P. blanchetiana , endemic to eastern and southeastern near-coastal Brazil; and P. confertiflora , with a bimodal distribution in northern French Guiana and northern Brazil. The Guianensis species complex possesses yellow to orange corollas, sepals glabrous and scarcely longer than broad to broader than long (distinctly longer than broad in P. longifolia ), inflorescences short- to longpedunculate, elongate, and laxly flowered, with deep yellow, yellow-orange or orange corollas ( P. grandiflora , P. longifolia , and some P. guianensis ). The Guianensis group comprises P. grandiflora , ranging over the Amazonian basin; P. guianensis , widely distributed over much of northern South America; and P. longifolia , in Venezuela, Guyana, and Surinam. Some specimens of Paypayrola guianensis have a shorter, moderately densely flowered inflorescence with the axis still visible, and corollas bright yellow rather than yellow-orange or orange, suggesting that it may itself represent a species complex with multiple undescribed taxa. The Hulkiana species complex possesses bright yellow corollas in a subsessile to short-pedunculate, moderately laxly to densely flowered inflorescence, and sepals puberulent and as long as to distinctly longer than broad. This last species complex consists of P. hulkiana , with a distribution in southeastern Venezuela east to French Guiana and south into northern Brazil, and P. panamensis , endemic to central Panama.

Comments:— Paypayrola panamensis is as distinct in features of its foliage, inflorescence, and floral characteristics as any other previously described species in the genus. The new species is similar to P. guianensis and P. hulkiana in certain traits but differs consistently in several non-overlapping qualitative and quantitative features as well as additional divergent tendencies. It is recognized here as a distinct and geographically disjunct Paypayrola species endemic to Panama. It is one of two species belonging to the Hulkiana species complex, and is the first described from Mesoamerica.

OTHER SPECIMENS EXAMINED: PANAMA. Coclé: La Pintada, Parque Nacional G.D. [General de Division] Omar Torrijos H. Rio San Juan [8°43’33” N, 80°39’57” W], 198 m, 22 July 2013, Martínez, Arcia, Rodríguez & Espinosa 1338 ( MO6468115 !) . La Pintada, Parque Nacional General de Division Omar Torrijos Herrera, La Rica [8°42’59” N, 80°35’30” W], 240 m, 23 July 2013, Ortiz, Zapata , Flores & Mora 1475 ( MO6614119 !, PMA) . Colón: Donoso Distrito, [Campamento de] Botija, Carretera pionera, 08°50’06”N 080°39’17”W, 11 September 2012, van der Werff, Gray, Martínez & Sanchez 24441 ( MO6450909 !, NY02714036 image!, US 3667240!) . Distrito de Donoso, [Campamento de] Botija, antigua cantera, sección 35, Cordenadas : N 8°50’00.8” W 80°38’59.8”, 181 m. altura, 16 January 2016, Batista 1512 , González & Lester ( NY04080735 image!) . MPSA Concession, Botija Camp, Along Quebrada Botija , 08°50’10”N 080°39’00”W, 22 May 2012, Hammel, De Gracia, Flores, Martínez, Merello & Quiel 26313 ( MO6450910 !) . Teck Cominco Petaquilla mining concession, along exploration road, 08°50’22”N 080°38’51”W, 184 m, 15 September 2007, McPherson 19558 ( MO6284425 !) . Teck Cominco Petaquilla mining concession, near transect C002 [8°50’22” N, 80°38’51” W], 205 m, 18 September 2007, McPherson 19679 ( MO6284424 !) . Teck Cominco Petaquilla mining concession, along road [8°49’29” N, 80°40’12” W], 323 m, 1 December 2007, McPherson & van der Werff 19910 ( MO6284426 !) . Teck Cominco Petaquilla mining concession, along ridge road [8°49’22” N, 80°39’32” W], 300 m, 25 February 2008, McPherson & Merello 20241 ( MO6284427 !) . Site of proposed copper mine (MPSA) [8°48’27” N, 80°36’20” W], 100 m, 13 December 2009, McPherson & Merello 21283 ( MO6208597 !) . Cocle del Norte , area del helipad BL02, tomando hacia el norte [8°51’47” N, 80°48’29” W], 20 July 2012, Aranda B. 4265 ( PMA) . Cocle del Norte , area del helipad CR10 , tomando hacia el sur [8°56’18” N, 80°41’19” W], 18 July 2012, Aranda B. 4233 ( PMA) . Río Taindi (Taimdi of maps) 6 km above confluence with Río Mandinga , 9°25’N, 79°11’W, 30–100 m, 5 April 1986, de Nevers & Herrera 7665 ( U1757196 !, WIS!) . Guna Yala. Between Rio Diablo & Rio Acuati near Nargana [mapped to 9°26’24”N, 78°34’12”W, same as other Duke collections in the area], 3 November 1967, Duke 14895 ( MO2191599 !) . 9°24’N, 78°34’W, 65 m, [no date], Galdames, Montenegro, Chung & Herrera 1587 ( MO5557973 !) . Trail to Cerro Óbu ( Habu of maps) from Río Urgandi ( Río Sidra ), 9°23’N, 78°48’W, 100–300 m, 24 June 1986, de Nevers, Herrera & Gernado 7974 ( MO04633270 !, U1757193 !) . Vicinidad de Río Diablo, desde el campamento Duque Sui hasta Isper Yala , 9°22’N, 78°35’W, 70–100 m, 3 July 1992, Herrera, B. Obaldía, R. Obaldía & Blanco 1210 ( MO5770374 !, WIS!) . 12 mi on Carti Road from Inter-American Highway then 3 hr walk along road down Atlantic side [mapped at 33 km from Delta Llano Carti at Pan-American Highway, 9°21’27” N, 78°58’40” W], 304 m, 15 February 1980, Antonio 3782 ( MO2936115 !) . Río Cangandí, hills W of river S of confluence with Río Titamibe , 9°24’N, 79°8’W, 50–150 m, 27 January 1985, de Nevers, Herrera, Charnley & Paredes 4669 ( MO3244722 !) . Río Cangandi at confluence of Quebrada Titamibe , 9°24’N, 79°7’W, 60 m, 8 February 1986, de Nevers & Herrera 7025 ( MO04633271 !, U1757198 !) . 16 km above Pan-Am Highway on road from El Llano to Carti-Tupile , Near campsite [9°19’08” N, 78°59’49” W], 350–400 m, 21 February 1973, Kennedy 2542a ( MO2906175 !, U1757199 !) . North of El Llano [9°19’05” N, 78°59’35” W], 13 February 1973, Dressler 4270 ( MO2153862 !) . Road from El Llano to Carti , 14.8 km north of the Panamerican Highway [9°19’03” N, 78°59’17” W], 300–500 m, 3 September 1977, Folsom & Maas 5221 ( MO2936070 !) . El Llano-Carthi Road , km 15 [9°19’05” N, 78°59’23” W], 300–400 m, 3 September 1977, Maas, Berg & Foulsen 2794 ( MO2639559 !, NY01340902 image!, U1757192 !) . Panamá: Along Llano-Carti road, c. 9°15’N, 79°00’W, 400 m, 24 November 1985, McPherson 7586 ( MO3311495 !) . Along Llano-Carti road, c. 9°15’N, 79°00’W, 400 m, 24 November 1985, McPherson 7588 ( MO3311494 !) . On El Llano-Carti road, near Nusigandi , along trail to waterfall, 9°15’N, 79°00’W, 250 m, 1 November 1992, McPherson & Richardson 16007 ( MO4251239 !) . El Llano-Cartí Road, Km 26.5, 9°19’N, 78°55’W, 200 m, 17 June 1986, de Nevers, Herrera, McPherson, D’Arcy & Allen 7828 ( MO04633269 !, U1757194 !) . El Llano-Cartí road, Km 22, 9°19’N, 78°55’W, 350 m, 17 June 1986, de Nevers & Herrera 7853 ( U1757195 !) . El Llano-Cartí Road, Km. 26.5, Trail NE from road, 9°19’N, 78°55’W, 175 m, 9 March 1985, de Nevers, Herrera & Charnley 5078 ( MO3244726 !) . El Llano-Cartí Road, Km. 26.5, trail to Río Cartí Chico and up ridge on E side, 9°19’N, 78°55’W, 200 m, 7 April 1985, de Nevers, Schatz, Herrera & Charnley 5236 ( MO3474324 !) . Nusagandi, Sede de Campo de PEMASKY, Approx. 20 km on El Llano-Carti Road , Trails near station, Aprox. 78°59’W, 9°18’N, 350 m, 29 March 1992, Paredes, Foster, Peña & Welton 657 ( F!) . El Llano-Cartí highway, 14–17 km. [mapped at 15.5 km along road, 9°19’5”N, 78°59’35”W] ca. 1/2 [way] between the Continental Divide and Atlantic coast opposite Carti, Near Atlantic base camp, 23 February 1973, Kennedy 2616 ( MO2985556 !) .

Figures

FIGURE 1. Features in life of flowering Paypayrola panamensis. A. Branch with leaves and inflorescences. B. Closeup of flowers. Photos by Barry Hammel.

FIGURE 2. Features of Paypayrola panamensis. A. Branches and leaves (de Nevers et al. 5363, holotype, MO4998611). B. Inflorescence, unbranched botryoid (de Nevers et al. 5363, isotype, WIS). C. Inflorescence, branched botryoid (McPherson 20241, MO6284427). D. Sepals (de Nevers et al. 5363, isotype, WIS). E. Petals; broadest (b) is conduplicate and folded in half (Maas et al. 2794, MO2639559). F. Intact reproductive structures (McPherson 7586, MO3311495). G. Androecium (de Nevers et al. 5363, isotype, WIS). H. Gynoecium (Maas et al. 2794, MO2639559). I. Capsule (McPherson 20241, MO6284427). J. Seed (McPherson 20241, MO6284427).

FIGURE 3. Map of the distribution of Paypayrola panamensis.